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Insight on the theropod fauna from the Uberaba Formation (Bauru Group),
Minas Gerais State: New megaraptoran specimen from the Late Cretaceous of
Brazil

Article in Rivista Italiana di Paleontologia e Stratigrafia · July 2013

DOI: 10.13130/2039-4942/6035

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 Rivista Italiana di Paleontologia e Stratigrafia volume 119 no. 2 pp. 205-214 July 2013

INSIGHT ON THE THEROPOD FAUNA FROM THE UBERABA FORMATION

(BAURU GROUP), MINAS GERAIS STATE: NEW MEGARAPTORAN SPECIMEN
FROM THE LATE CRETACEOUS OF BRAZIL

AGUSTIÂN G. MARTINELLI 1 *, LUIZ CARLOS BORGES RIBEIRO 1, ARIEL H. MEÂNDEZ 2,

FRANCISCO MACEDO NETO 1, CAMILA LOURENCINI CAVELLANI 1, EDUARDO FELIX 1,
MARA LUÂCIA DA FONSECA FERRAZ 1 & VICENTE DE PAULA ANTUNES TEIXEIRA 1
Received: January 1st 2013; accepted: February 21, 2013

Key words: Uberaba Formation, Megaraptora, caudal vertebra, ulteriori informazioni sull'anatomia della coda di questo bizzarro grup-
pneumaticity. po.
Abstract. The first bony theropod record from the Campanian
Uberaba Formation (Bauru Group) is described. It consists of an iso-
lated caudal centrum (CPPLIP 1324) found in the city of Uberaba, Introduction

Minas Gerais State, Brazil. The amphicoelous centrum possesses a

length to height ratio of 1.74, deep elliptical lateral pneumatic foramen The Late Cretaceous continental vertebrate as-
representing 26% of centrum length with three main sub-circular air semblages from Brazil have been focus of systematic
chambers, and camellate internal structure. This combination of fea- researches in the last decades, considerably increasing
tures is shared with Aerosteon, Megaraptor, and Orkoraptor from the their fossiliferous content (e.g., Kellner & Campos
Late Cretaceous of Argentina and with the Megaraptora indet. from the 2000; Bittencourt & Langer 2011). Particularly, rocks
SaÄo Jose do Rio Preto Formation (Bauru Group), SaÄo Paulo State, from the Bauru Group outcropping in southeastern
allowing us to refer it to the Megaraptora clade (Tetanurae, Neovena- Brazil have provided the most abundant and taxonomi-
toridae). As such, the new specimen represents the second megaraptor-
an from the Late Cretaceous of Brazil and provides new information on cally varied dinosaur fauna from the Upper Cretaceous
tail anatomy on this bizarre group. of this country (Bittencourt & Langer 2011). Titanosaur
sauropods are known to occur in most of all the recog-
Riassunto. Viene qui descritto il primo ritrovamento di ossa di nized formations of the Bauru Group (e.g., Santucci &
teropode dalla Formazione Uberaba (Gruppo Bauru) di etaÁ Campania- Bertini 2001; Kellner et al. 2006; Santucci 2008; Salgado
na. Si tratta di un corpo vertebrale caudale isolato (CPPLIP 1324) & Carvalho 2008), commonly including associated par-
trovato nella cittaÁ di Uberaba, Stato di Minas Gerais, Brasile. Il corpo
vertebrale anficele ha rapporto lunghezza/altezza di 1.74, un profondo tial remains. In contrast, the theropod record is far less
foramen pneumatico laterale ellittico che rappresenta il 26% della lun- abundant by means of isolated occurrence (teeth and
ghezza del corpo con tre cavitaÁ pneumatiche principali di forma sub- partial bones) in several localities of the Bauru Group
circolare, e struttura interna a nido d'ape. Questa combinazione di (e.g., Kellner & Campos 2002; Novas et al. 2005, 2008;
caratteri eÁ condivisa con Aerosteon, Megaraptor e Orkoraptor del Cre- Bittencourt & Langer 2011).
taceo superiore dell'Argentina e con il Megaraptora indet. dalla Forma- Theropod dinosaurs from the Cretaceous of Bra-
zione di SaÄo Jose do Rio Preto (Gruppo Bauru), dello Stato di San
Paolo, consentendo di riferire il reperto al clade Megaraptora (Tetanu- zil are still inadequate known; nonetheless, their diver-
rae, Neovenatoridae). In quanto tale, il nuovo esemplare rappresenta il sity has been increased considerably in the last decades
secondo Megaraptora dal Cretaceo Superiore del Brasile e fornisce (see below). Pre-Santonian non-avian theropods in-

1 Centro de Pesquisas PaleontoloÂgicas L. I. Price, Complexo Cultural e CientõÂfico PeiroÂpolis (CCCP/UFTM), BR-262, Km 784, Bairro
PeiroÂpolis, 38039-755, Uberaba, Minas Gerais, Brazil.
2 CONICET-Instituto de Investigaciones en Biodiversidad y Medioambiente, Quintral 1250, Bariloche, RõÂo Negro, Argentina. MPB
(Museo PaleontoloÂgico Bariloche), Av. 12 de Octubre y Sarmiento, Bariloche, RõÂo Negro, Argentina.
* Corresponding author: agustõÂ[email protected]
206 Martinelli A.G. et al.

clude: a) teeth of noasaurids (Lindoso et al. 2012); b) a precise level is unknown, the green fine sandstone that
teeth and few vertebral elements of carcharodontosaur- covered the bone would likely correspond to the layers
ids (Vilas BoÃas et al. 1999; Medeiros & Schultz 2002; of the upper portion of the unit. A great portion of the
Medeiros et al. 2007); c) the spinosaurids Irritator chal- Uberaba city was constructed over this unit which is
lengeri (Martill et al. 1996), Angaturama limai (Kellner restricted to this portion of the TriaÃngulo Mineiro re-
& Campos 1996; it is considered by some authors as a gion (Hasui 1968; Fernandes & Coimbra 2000; Batezelli
junior synonym of I. challenger; e.g., Sereno et al. 1998; et al. 2007). Relatively small outcrops can be seen along
Sues et al. 2002), Oxalaia quilombesis (Kellner et al. the city, in road and train rail cuts, in parks or aban-
2011), and isolated spinosaurid teeth and bones (Cam- doned quarries. The Uberaba Formation is laying in
pos & Kellner 1991; Kellner & Campos 1999; Medeiros discordance with the basaltic rocks of the Serra Geral
& Schultz 2002; Bittencourt & Kellner 2004; Machado Formation, and at the top, only observed in a few
& Kellner 2005, 2007; Medeiros 2006); d) the basal coe- places, it is covered by the MarõÂlia Formation (Fer-
lurosaurian Santanaraptor placidus (Kellner 1999; No- nandes & Coimbra 2000). To the northwest, the Uber-
vas et al. 2012b); and e) the compsognathid Mirischia aba Formation has a roughly lateral contact with the
asymmetrica (Naish et al. 2004). On the other hand, Adamantina Formation (Goldberg & Garcia 1995),
post-Santonian theropod record includes abelisaurids and most stratigraphic columns of the Bauru Basin
(e.g., Bertini 1996; Kellner & Campos 2002; Novas et placed Uberaba and Adamantina (Vale do Rio do Peixe
al. 2008; Candeiro et al. 2012a), megaraptorans (MeÂndez Formation in Fernandes & Coimbra 2000) as laterally
et al. 2012), and maniraptorans (Bertini et al. 1997; Ber- correlated formations (e.g., Fernandes & Coimbra 2000;
tini & Franco-Rosas 2001; Novas et al. 2005; Machado Batezelli et al. 2007).
et al. 2008; Candeiro et al. 2012b). Scanty post-Santo- The Uberaba Formation comprises a continental
nian carcharodontosaurid remains were also reported sequence of reddish siltstones interbedded with greenish
from a few localities of Minas Gerais and SaÄo Paulo in massive sandstones, with conglomeratic lenses, espe-
rocks from the Bauru Group (e.g., Candeiro et al. 2006, cially at the base (Fernandes & Coimbra 2000), devel-
2012a; Azevedo et al. 2013); nonetheless, most records oped in a braided fluvial system (Ferreira JuÂnior &
are likely dubious, pending further studies (see Canale Guerra 1995; Fernandes & Coimbra 2000). The age of
et al. 2009; Brusatte et al. 2007; Souza et al. 2011, among this formation is still out of consensus due to it scanty
others). The only formally named theropod and hither- fossil record and indirect inferences (see Santucci 2008).
to one of the most complete from post-Santonian times Gobbo-Rodrigues et al. (1999) proposed a Cam-
of Brazil is the abelisaurid Pycnonemosaurus nevesi, panian-Maastrichtian age for the Adamantina Forma-
from the Parecis Group of Mato Grosso State (Kellner tion based on ostracods from the SaÄo Paulo State. Con-
& Campos 2002). All other theropod records are recog- sidering the lateral correlation, a similar age for the
nized on the basis of isolated elements. Uberaba Formation at Minas Gerais State has been as-
In this contribution, a partial caudal vertebra sumed (Santucci 2008). Magnetostratigraphic studies of
(CPPLIP 1324) referred to the theropod clade Mega- the Uberaba and MarõÂlia Formations (Tamrat et al.
raptora is described and compared. It constitutes the 2002) indicate that the deposition of Uberaba could
second megaraptoran record from the Upper Creta- not be older than Campanian; therefore, this data cor-
ceous of Brazil and the first theropod bone from the roborated a post-Santonian age. In contrast, Dias-Brito
Uberaba Formation. Until now, the sparse fossil record et al. (2001), also based on ostracods, considered a San-
of the Uberaba Formation (Bauru Group) was limited tonian age for the top of the Adamantina Formation,
to titanosaur sauropods (Santucci 2008) and three asso- and by inference it was used for the Uberaba Forma-
ciated eggs, briefly described, referred to Theropoda tion. Moreover, these authors noted a depositional hia-
(Kellner et al. 1998; Kellner & Campos 2000). tus of 11 Ma between Adamantina and MarõÂlia Forma-
tions (Dias-Brito et al. 2001). Recently, Montefeltro et
al. (2011) discussed the age of the outcrops of the Ada-
mantina Formation (= Vale do Rio do Peixe Formation)
at west TriaÃngulo Mineiro and based upon baurusuchid
Geological Settings

The specimen here reported was found in 2011 crocodyliforms suggested biostratigraphic differences
during the construction of the Uberaba Regional Hos- between the outcrops from SaÄo Paulo and Minas Gerais
pital building (``Hospital Regional de Uberaba'') located states. Pissarrachampsa sera from west TriaÃngulo Mine-
in front of the SaÄo JoaÄo Batista Cementery, Uberaba iro, and also Campinasuchus dinizi from the same unit
city (Minas Gerais State, Brazil), without possibility and found in a near locality (Carvalho et al. 2011), are
of access to the outcrop. Based on the rock surrounding closer relatives to Wargosuchus australis from the San-
the bone before preparation and the location, it cer- tonian Bajo de la Carpa Formation (NeuqueÂn Group,
tainly comes from the Uberaba Formation. Although Argentina) rather than to baurusuchines, which are re-
 New megaraptoran specimen from the Late Cretaceous of Brazil 207

stricted to the Adamantina Formation outcropping at Systematic paleontology

SaÄo Paulo State (Montefeltro et al. 2011). Therefore, Theropoda Marsh, 1881
the data based on baurusuchids is not conclusive (Mon- Tetanurae Gauthier, 1986
tefeltro et al. 2011) but highlights the difficult to corre-
late units based on tetrapod record for age interpreta- Neovenatoridae Benson et al., 2010
tions. Because the correlation of the outcrops of the Megaraptora Benson et al., 2010
Adamantina Formation of Minas Gerais and SaÄo Paulo Gen. et sp. indet., fig.1-2
has not been fully explored (e.g., Montefeltro et al.
2011) and due to the broad distribution of this unit, Referred Material: CPPLIP 1324, middle caudal vertebral cen-
differences in age and lithofacies are expected. trum (Fig. 1, 2).
Locality, horizon and age: CPPLIP 1324 was found during the
The proposal of a Campanian-Maastrichtian age construction of the building of the Uberaba Regional Hospital located
for the Uberaba Formation is problematic because the in front of the Uberaba Cemetery (GPS: 19ë44'29.41"S/
supposed coetaneity of Adamantina/Uberaba forma- 47ë57'34.96"W), Uberaba city, Uberaba County, Minas Gerais State,
tions, at least in part, and the partial coetaneity with Brazil; Uberaba Formation, Bauru Group, Campanian, Upper Cretac-
eous (see Geological Settings).
the youngest and overlaying MarõÂlia Formation (Maas-
trichtian). A pre-Campanian age for the Uberaba For- Description. CPPLIP 1324 consists of an isolated
mation contradicts magnetostratigraphic studies (Tam- middle caudal vertebral centrum (Fig. 1) of a medium-
rat et al. 2002); therefore, (a) the Uberaba Formation has sized theropod dinosaur. The centrum is almost com-
not a total lateral correlation with the Adamantina For- plete with the outline of the articular surfaces eroded,
mation or (b) the age based on ostracodes from the being the posterior edge more damaged. A remnant of
Adamantina Formation is not conclusive for inferring transverse process is partially preserved on the right side
the age of the Uberaba Formation. (Fig. 1). The centrum measurements are 8 cm maximum
Considering the faunal record of the younger long, 4.6 cm maximum tall and 6.1 cm maximum width.
Maastrichtian MarõÂlia Formation at TriaÃngulo Mineiro Its general aspect is anteroposteriorly long and dorso-
region (e.g., Bertini et al. 1993; Santucci & Bertini 2001; ventrally short. The length to height (L/h) ratio is 1.74.
Salgado & Carvalho 2008), the above mentioned issues CPPLIP 1324 is interpreted as a middle caudal centrum
(e.g., magnetostratigraphy), and due to the large span of because it is anteroposteriorly elongated, and transver-
the interval Campanian-Maastrichtian proposed for the sely wider than high. In Neovenator anterior vertebral
Uberaba Formation, we prefer to use a restricted Cam- caudal bodies are higher dorsoventrally than wide trans-
panian age for the Uberaba Formation, a matter which versely (Brusatte et al. 2008). In more posterior ele-
should be accurately addressed in future contributions. ments, starting at the ?fourteenth caudal (caudal G),

Fig. 1 - Megaraptora indet. from the Uberaba Formation (Bauru Group), Uberaba, Minas Gerais State, Brazil. CPPLIP 1324, posterior caudal
centrum in: A) left lateral, B) right lateral, C) anterior, D) dorsal, E) ventral, and F) posterior views. Scale bar equals 20 mm.
208 Martinelli A.G. et al.

the articular surfaces are wider than high (Brusatte et al. centrum is dominated by a deep elliptical shaped pneu-
2008). The transverse process is also high in anterior matic foramen that represents 26% of the centrum
caudals and in the ?fourteenth caudal (caudal G) it is length (Fig. 1, 2). It is located at middle height and
placed in a lower position (Brusatte et al. 2008). Based length. On the right side, the pneumatic foramen is
upon these features, we likely positioned the specimen shallower, with three main sub-circular air chambers
CPPLIP 1324 as a middle caudal body, posterior to the separated by thin bone laminae. The anterior and pos-
caudal G of Neovenator (Brusatte et al. 2008). terior chambers are similar in size, about three times
The centrum is amphicoelous with the articular smaller than the central one. Near the anterior edge of
surfaces sub-circular in outline, except for the dorsal the pneumatic foramen, there are very small foramina,
margin that forms the floor of the neural canal, which possibly nutritious ones. On the left side, the pneumatic
is concave. The articular surfaces have broad sulci in- foramen is deeper, with at least one thin lamina of bone
dicating attachment for cartilaginous tissues. The edges separating two main air chambers. The anterior one is
of the articular surfaces are broken, showing a strongly the largest but probably it constitutes two collapsed
camellate internal tissue (Fig. 2). The camellate structure chambers, with the separating wall broken off.
is constituted of chambers that are subrectangular in Tomography of CPPLIP 1324 reveals the changes
cross-section, separated one to each other by thin lami- in size of the internal chambers and the connections of
nae of bone. The chambers exhibit a regular placement the chambers of the pneumatic foramina in the center of
along the articular edge and are larger at the ventral the vertebral body (Fig. 2). At the ˆ anterior and pos-
margin of the articular surfaces and at the conjunction terior portions of the centrum, the internal chambers are
with the neural arch. In lateral view, each side of the larger near the edge of the bone forming a periphery

Fig. 2 - Details of the pneumatic foramen and camellate internal structure of Megaraptora indet. CPPLIP 1324. Caudal centrum in right
lateral.
 New megaraptoran specimen from the Late Cretaceous of Brazil 209

band, and become smaller to the inner side. Moving to Novas et al. 2008; Calvo et al. 2004) from the Late
the longitudinal center, the peripheral chambers become Cretaceous of Argentina are the only South American
even larger. At the portion of the pneumatic foramen, named theropods with pneumatic foramen on caudal
chambers are notably great with slightly broader bone centra. They were recently nested into the Megaraptora
laminae. It can be seen that the chambers of the pneu- clade, within a large group, called Carcharodontosauria
matic foramina open internally and contact one to each (see Benson et al. 2010 and Carrano et al. 2012; although
other (Fig. 2). coelurosaurian affinities were proposed recently, see
In dorsal view, the centrum has a relatively broad, Novas et al. 2012). The size and depth of pneumatic
transversely concave floor of the neural canal, bearing a foramen of CPPLIP 1324 are more similar to Aerosteon
pair of small, elliptic foramina just posterior to the (Fig. 3) than to other megaraptorans. In Aerosteon, the
transversal middle line of the centrum (Fig. 1). The re- middle centrum is proportionally shorter anteroposter-
gion for the attachment of the neural arch occupies most iorly than in CPPLIP 1324, possibly because the Brazi-
of the centrum length, leaving only a small portion free lian specimen occupies a slightly posterior position in
at the front. On the right side, there is a small portion of the tail. Despite the fragmentary nature of CPPLIP
transverse process without well defined edges. The 1324, we refer it to the neovenatorid clade Megaraptora
transverse process is in a low position and it is highly due to the large pneumatic foramen and the camellate
pneumatic as the centrum. It is tightly attached to the internal structure.
body without any evidence of suture, indicating that According to Brussate et al. (2008), the pneumatic
this element belongs to a mature individual. Moreover, foramina are prominent in anterior caudals of the basal
the surface at the left side where the base of the neural neovenatorid Neovenator but posteriorly they are shal-
arch should be placed was covered by rock suggesting lower and poorly expressed, being absent up to the
that it was broken before burial. ?twenty first element (caudal J). According to the pro-
The ventral surface is almost flat with concave portions of the centrum, the position of the base of the
lateral outlines when observed in ventral view. On the transverse process, and the almost flat ventral surface,
axial line, there is a very shallow elevation that does not the presence of such a large pneumatic foramen in the
reach to constitute a real crest. Because ventral edges of lateral surface of CPPLIP 1324, in that portion of the
the articular surfaces are eroded, there is no clear evi- tail, is likely a difference with Neovenator (Fig. 3A).
dence of facets for haemal arch (Fig. 1). Possibly, these Although preliminary, it can be indicating that mega-
facets are not evident because they usually are less visi- raptorans have a more advance pneumatic condition
ble in middle and posterior caudals, due to the reduc- than in the basal Neovenator. Furthermore, the Brazi-
tion of the haemal arches. lian specimen differs from Neovenator (Brusatte et al.
2008; Fig. 3A) because the base of the transverse process
is located slightly posterior in comparison to the British
theropod.
CPPLIP 1324 can be excluded from carcharodon-
Discussion

Taxonomical assignment of CPPLIP 1324. Among tosaurids, particularly those coming from Patagonia
the theropod record from Brazil, CPPLIP 1324 exhibits (e.g., Mapusaurus, Giganotosaurus), because in the latter
close resemblances with the recently described Mega- the caudals lack real pneumatic foramen (e.g., Coria &
raptora gen. et sp. indet. (MPMA 08-003-94) from the Currie 2006). An anterior caudal assigned to Carcharo-
SaÄo Jose do Rio Preto Formation (MeÂndez et al. 2012), dontosaurus by Stromer (1931), from the early Late
found at IbiraÂ, SaÄo Paulo State, Brazil (Fig. 3C). As in Cretaceous of Africa, has a pneumatic foramen, differ-
MPMA 08-003-94, the caudal centrum from Uberaba ent to the condition of South American carcharodonto-
has notorious pneumaticity evidenced by a large lateral saurids. Direct comparison with CPPLIP 1324 cannot
pneumatic foramen and camellate internal structure (see be done because they have different position on the tail.
Fig. 2). Nonetheless, the resemblance of CPPLIP 1324 with
Among theropods, the caudal vertebral pneuma- other megaraptorans (e.g., Aerosteon) than with most
ticity occurred independently among distinctive clades carcharodontosaurids is notorious.
such as ceratosaurs, megaraptorans, carcharodontosaur- Due to the pneumatic condition of CPPLIP 1324,
ids, and some coelurosaurians (e.g., oviraptorosaurians comparison with therizinosauroids and oviraptorosaurs
and therizinosaurians) (Stromer 1931; Britt 1993; Sues are pertinent. Nonetheless, material of the first group
1997; Xu et al. 2002; O'Connor & Claessens 2005; has not been documented so far in South America and
O'Connor 2006; Barsbold et al. 2000; Brusatte et al. the occurrence of oviraptorosaurs in South America was
2008; Novas et al. 2008; Sereno et al. 2008; Zanno et dismissed (see Agnolin & Martinelli 2007). As in the
al. 2009; Benson et al. 2010, 2012). Particularly, Aero- Brazilian specimen, oviraptorosaurs and most therizino-
steon, Megaraptor, and Orkoraptor (Sereno et al. 2008; sauroids have caudal centra with a pneumatic foramen
210 Martinelli A.G. et al.

Fig. 3 - Comparison of caudal vertebrae among Neovenatoridae in left lateral, ventral and dorsal views. A) Neovenator salerii, caudal vertebra
``I'' (modified from Brusatte et al. 2008: Plate 19); B) Megaraptora indet. CPPLIP 1324; C) Megaraptora indet. MPMA 08-003-94
(lateral and ventral views modified from MeÂndez et al. 2012); D) Aerosteon riocoloradensis, middle caudal centrum (modified from
Sereno et al. 2008 and MeÂndez et al. 2012). Scale bar equals 50 mm.

that is inter- and intra-specifically variable (e.g., Sues respectively). A very subtle median elevation on the
1997; Barsbold et al. 2000; Zhang et al. 2001; OsmoÂlska ventral surface of CPPLIP 1324 is observed, being to-
et al. 2004; Xu et al. 2002, 2007; Zanno et al. 2009). Differ- tally absent in the Ibira specimen. These differences
ing from CPPLIP 1324, the pneumatic foramen in these would correspond to different position in the tail, being
groups is positioned very close to the transverse process CPPLIP 1324 anterioriorly positioned on the tail than
(e.g., Sues 1997; Xu et al. 2007) and it is proportionally MPMA 08-003-94. Although the difference in the tail
smaller than in CPPLIP 1324. Furthermore, oviraptoro- position, it is noteworthy that MPMA 08-003-94 is ap-
saurs usually have caudals with a medial groove (shal- proximately 30% larger than CPPLIP 1324 indicating a
low or deep) on the ventral surface of the centrum, larger individual/species in the Maastrichtian of SaÄo
delimited by longitudinal ridges (e.g., Sues 1997; Xu Paulo. Despite specimens are isolated they share close
et al. 2007), differing from the condition of CPPLIP resemblances supporting their phylogenetic proximity.
1324. Although the comparison is significantly limited, Both individuals come from different units from the
the absent of these coelurosaurians in the South Amer- Bauru Group. CPPLIP 1324 is considered to be Cam-
ican fossil record alert us on the referral of the Brazilian panian in age (see Geological Settings), from the Ube-
specimen to these groups, also taking into consideration raba Formation, and MPMA 08-003-94 to be Maastrich-
the isolated and fragmentary nature of CPPLIP 1324. tian, coming from the SaÄo Jose do Rio Preto Formation
(Bertini & Menegazzo 2009; MeÂndez et al. 2012). They
Comparisons with the other Brazilian megarap- clearly show the occurrence of still poorly known ther-
toran. In general aspect, CPPLIP 1324 and MPMA opod lineages in Brazil, and MPMA 08-003-94 repre-
08-003-94 (MeÂndez et al. 2012) are quite similar. The sents the youngest record of this clade, together with
main differences are that the pneumatic foramen of Orkoraptor from the Maastrichtian Pari Aike Forma-
CPPLIP 1324 is deeper, located at mid height and tion of Patagonia, Argentina (Novas et al. 2008).
length of the body of the centrum, and with discrete
edges. In MPMA 08-003-94 the pneumatic foramen,
located in a slightly higher position, slopes gradually
the lateral surface, and the internal divisions are not
Conclusion

symmetrical (Fig. 3C). In CPPLIP 1324 the L/h ratio CPPLIP 1324 represents the second Megaraptor-
is slightly lower than in MPMA 08-003-94 (1.7 vs. 1.9, an record from Brazil, being both occurrences confined
 New megaraptoran specimen from the Late Cretaceous of Brazil 211

to the Late Cretaceous of the Bauru Group. With the terrestrial and hyper-carnivorous adaptations (e.g., Gas-
exception of Pycnonemosaurus nevesi, from the Parecis parini et al. 1993; Candeiro & Martinelli 2006; Riff &
Group of Mato Grosso State (Kellner & Campos Kellner 2011), we believe that pertinent morph-spatial
2002), the theropod record from Bauru Group is still and morph-ecological comparisons in detail are still
poorly represented, based upon fragmentary and iso- poorly explored. In contrast, the theropod record is
lated material (teeth, vertebrae, claw, furcula, etc.) that growing both taxonomically and in number of available
allow limited taxonomic analyses, and some records specimens. As such, the inferences on the supposed taxa
must be carefully evaluated. The new specimen also that occupied the terrestrial carnivorous niches are
represents the first theropod bone from the Uberaba likely preliminary and still unrealistic.
Formation, which until now only yielded titanosaur
remains (Santucci 2008) and theropod eggs (Kellner et Acknowledgements. We thank the financial support for this
al. 1998). research provided by the FundacËaÄo de Amparo aÁ Pesquisa do Estado
The recent findings of Megaraptorans in the SaÄo de Minas Gerais (FAPEMIG), the Conselho Nacional de Desenvolvi-
Jose do Rio Preto Formation (MeÂndez et al. 2012) and mento CientõÂfico e TecnoloÂgico (CNPq), and the FundacËaÄo de Ensino e
now in the Uberaba Formation (this paper) enrich the Pesquisa de Uberaba (FUNEPU). We especially thank S. Brusatte, F.V.
still poorly understood theropod fossil record in central Iori, and J.I. Canale because provided useful information on Neovena-
tor, Megaraptora indet. (MPMA 08-003-94), and Giganotosaurus, re-
South America by the Upper Cretaceous. Although it spectively, and R. Goulart due to tomography assistance at the UFTM
was briefly suggested that mesoeucrocodylians, such as (Uberaba). The reviewers S. Brusatte, S. Maganuco, F. Agnolin and the
baurusuchids, occupied the ecological niches of thero- Editor L. Rook provided useful comments that greatly improved the
pods due to their abundant fossil record and several Ms.

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